British spy Finn McMissile infiltrates the world's largest untapped oil reserves, owned by a group of "lemon" cars. After being discovered, he is forced to flee and fake his death.
Four-time Piston Cup champion Lightning McQueen returns home to Radiator Springs and reunites with his best friend Mater and his girlfriend Sally Carrera. Former oil tycoon Sir Miles Axlerod, now a green power advocate, announces a racing series called the "World Grand Prix" to promote Allinol, his biofuel. When Italian formula race car Francesco Bernoulli challenges McQueen, he and Mater - along with Luigi, Guido, Fillmore, and Sarge - depart for Tokyo, Japan for the World Grand Prix.
Meanwhile, the lemons, led by Professor Zündapp and an unknown mastermind, secretly plot to secure their oil profits by using a weapon disguised as a television camera to ignite the Allinol fuel. McMissile and partner Holley Shiftwell attempt to rendezvous with American spy car Rod "Torque" Redline at a World Grand Prix promotional event in Tokyo, to receive information about the mastermind; however, Redline is attacked by Zündapp's henchmen and passes his information to Mater before he is captured. Holley and Finn mistake Mater as their American contact. Zündapp tortures Torque and reveals that Allinol can ignite if impacted by a high electromagnetic pulse. Zündapp demonstrates it on Torque, killing him, but not before they realize that he passed this information to Mater.
During the first race, Finn and Holley help Mater evade Zündapp's henchmen; in the process, Mater inadvertently gives McQueen bad racing advice which causes him to lose the race to Bernoulli. Meanwhile, Zündapp uses the weapon on several race cars. After McQueen falls out with Mater, Finn - who still believes Mater is an American spy - drafts him into foiling Zündapp's plot. Finn and Mater escape Zündapp's henchmen and climb aboard Siddeley. Finn and Holley remove the tracking device on Mater and discover in it a picture of a mysterious British engine, which Mater identified by the engine and rare parts.
Finn, Holley, and Mater fly to Paris, France, where they go into a black-market and meet an old friend of Finn's, Tomber. Finn and Holley show the mysterious engine to Tomber, who tells them that the car with the engine was his best customer, but he has never met him in person. Mater explains what he knows about the evil Lemons, and realizes that every Lemon involved with the plot is one of "history's biggest loser cars." Tomber tells Finn, Holley and Mater that the lemons are going to have a secret meeting in Porto Corsa, Italy (where the next race in the World Grand Prix is taken place).
In Italy, Mater infiltrates the criminals' meeting and discovers Zündapp's plan. Zündapp's henchmen, meanwhile, use their weapon on several more cars during the race, eventually causing a multi-car pileup on the Casino Bridge. With the Allinol fuel under suspicion, Axlerod suspends its use for the final race in England; however, McQueen chooses to continue using it. The criminals decide to kill McQueen in the next race; upon hearing this, Mater is exposed and is captured along with McMissile and Shiftwell, and tied up inside Big Bentley's bell tower in London, England.
Mater realizes how foolishly he has been acting. The criminals use the weapon on McQueen during the race, but nothing happens. Mater escapes to warn his friends of a bomb planted in McQueen's pit stop, but McMissile and Shiftwell find that the bomb was actually planted on Mater. They warn Mater about the bomb, and he flees to protect his friends. However, he is pursued by McQueen in an attempt to reconcile, unaware of the danger until they are beyond the range of Zündapp's remote detonator. He sends his henchmen to kill McQueen and Mater, but they are foiled by the combined efforts of McMissile, Shiftwell, and the Radiator Springs residents. Upon his capture, Zündapp reveals that only the person who installed the bomb can deactivate it, and Mater realizes that Axlerod is the mastermind behind the plot. Mater confronts Axlerod, trapping him next to the bomb. Axlerod finally deactivates the bomb, and he, Zündapp and the lemons are arrested by the police.
As a reward for his heroism, Mater receives an honorary knighthood from the Queen and returns home with his friends, where the cars from the Grand Prix take part in the unofficial Radiator Springs Grand Prix. Fillmore reveals that before the last race, Sarge replaced McQueen's Allinol with Fillmore's organic fuel, which prevented McQueen from being affected by the weapon. McMissile and Shiftwell invite Mater to join them in another spy mission, but he graciously turns it down but asks Shiftwell for a date when she returns which she accepts. He gets to keep the rockets they gave him earlier, which he uses in the Radiator Springs race.
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